The Variation of Animals and Plants under Domestication · Charles Darwin
REFERENCES
_On the Good derived from slight Changes in the Conditions of Life._—In considering whether any facts were known which might throw light on the conclusion arrived at in the last chapter, namely, that benefits ensue from crossing, and that it is a law of nature that all organic beings should occasionally cross, it appeared to me probable that the good derived from slight changes in the conditions of life, from being an analogous phenomenon, might serve this purpose. No two individuals, and still less no two varieties, are absolutely alike in constitution and structure; and when the germ of one is fertilised by the male element of another, we may believe that it is acted on in a somewhat similar manner as an individual when exposed to slightly changed conditions. Now, every one must have observed the remarkable influence on convalescents of a change of residence, and no medical man doubts the truth of this fact. Small farmers who hold but little land are convinced that their cattle derive great benefit from a change of pasture. In the case of plants, the evidence is strong that a great advantage is derived from exchanging seeds, tubers, bulbs, and cuttings from one soil or place to another as different as possible.
The belief that plants are thus benefited, whether or not well founded, has been firmly maintained from the time of Columella, who wrote shortly after the Christian era, to the present day; and it now prevails in England, France, and Germany.[1] A sagacious observer, Bradley, writing in 1724,[2] says, “When we once become Masters of a good Sort of Seed, we should at least put it into Two or Three Hands, where the Soils and Situations are as different as possible; and every Year the Parties should change with one another; by which Means, I find the Goodness of the Seed will be maintained for several Years. For Want of this Use many Farmers have failed in their Crops and been great Losers.” He then gives his own practical experience on this head. A modern writer[3] asserts, “Nothing can be more clearly established in agriculture than that the continual growth of any one variety in the same district makes it liable to deterioration either in quality or quantity.” Another writer states that he sowed close together in the same field two lots of wheat-seed, the product of the same original stock, one of which had been grown on the same land and the other at a distance, and the difference in favour of the crop from the latter seed was remarkable. A gentleman in Surrey who has long made it his business to raise wheat to sell for seed, and who has constantly realised in the market higher prices than others, assures me that he finds it indispensable continually to change his seed; and that for this purpose he keeps two farms differing much in soil and elevation.
With respect to the tubers of the potato, I find that at the present day the practice of exchanging sets is almost everywhere followed. The great growers of potatoes in Lancashire formerly used to get tubers from Scotland, but they found that “a change from the moss-lands, and _vice versa,_ was generally sufficient.” In former times in France the crop of potatoes in the Vosges had become reduced in the course of fifty or sixty years in the proportion from 120-150 to 30-40 bushels; and the famous Oberlin attributed the surprising good which he effected in large part to changing the sets.[4] A well-known practical gardener, Mr. Robson[5] positively states that he has himself witnessed decided advantage from obtaining bulbs of the onion, tubers of the potato, and various seeds, all of the same kind, from different soils and distant parts of England. He further states that with plants propagated by cuttings, as with the Pelargonium, and especially the Dahlia, manifest advantage is derived from getting plants of the same variety, which have been cultivated in another place; or, “where the extent of the place allows, to take cuttings from one description of soil to plant on another, so as to afford the change that seems so necessary to the well-being of the plants.” He maintains that after a time an exchange of this nature is “forced on the grower, whether he be prepared for it or not.” Similar remarks have been made by another excellent gardener, Mr. Fish, namely, that cuttings of the same variety of Calceolaria, which he obtained from a neighbour, “showed much greater vigour than some of his own that were treated in exactly the same manner,” and he attributed this solely to his own plants having become “to a certain extent worn out or tired of their quarters.” Something of this kind apparently occurs in grafting and budding fruit-trees; for, according to Mr. Abbey, grafts or buds generally take with greater facility on a distinct variety or even species, or on a stock previously grafted, than on stocks raised from seeds of the variety which is to be grafted; and he believes this cannot be altogether explained by the stocks in question being better adapted to the soil and climate of the place. It should, however, be added, that varieties grafted or budded on very distinct kinds, though they may take more readily and grow at first more vigorously than when grafted on closely allied stocks, afterwards often become unhealthy.
I have studied M. Tessier’s careful and elaborate experiments[6] made to disprove the common belief that good is derived from a change of seed; and he certainly shows that the same seed may with care be cultivated on the same farm (it is not stated whether on exactly the same soil) for ten consecutive years without loss. Another excellent observer, Colonel Le Couteur[7] has come to the same conclusion; but then he expressly adds, if the same seed be used, “that which is grown on land manured from the mixen one year becomes seed for land prepared with lime, and that again becomes seed for land dressed with ashes, then for land dressed with mixed manure, and so on.” But this in effect is a systematic exchange of seed, within the limits of the same farm.
On the whole the belief, which has long been held by many cultivators, that good follows from exchanging seed, tubers, etc., seems to be fairly well founded. It seems hardly credible that the advantage thus derived can be due to the seeds, especially if very small ones, obtaining in one soil some chemical element deficient in the other and in sufficient quantity to influence the whole after-growth of the plant. As plants after once germinating are fixed to the same spot, it might have been anticipated that they would show the good effects of a change more plainly than do animals which continually wander about; and this apparently is the case. Life depending on, or consisting in, an incessant play of the most complex forces, it would appear that their action is in some way stimulated by slight changes in the circumstances to which each organism is exposed. All forces throughout nature, as Mr. Herbert Spencer[8] remarks, tend towards an equilibrium, and for the life of each organism it is necessary that this tendency should be checked. These views and the foregoing facts probably throw light, on the one hand, on the good effects of crossing the breed, for the germ will be thus slightly modified or acted on by new forces; and on the other hand, on the evil effects of close interbreeding prolonged during many generations, during which the germ will be acted on by a male having almost identically the same constitution.
_Sterility from Changed Conditions of Life._ I will now attempt to show that animals and plants, when removed from their natural conditions, are often rendered in some degree infertile or completely barren; and this occurs even when the conditions have not been greatly changed. This conclusion is not necessarily opposed to that at which we have just arrived, namely, that lesser changes of other kinds are advantageous to organic beings. Our present subject is of some importance, from having an intimate connection with the causes of variability. Indirectly it perhaps bears on the sterility of species when crossed: for as, on the one hand, slight changes in the conditions of life are favourable to plants and animals, and the crossing of varieties adds to the size, vigour, and fertility of their offspring; so, on the other hand, certain other changes in the conditions of life cause sterility; and as this likewise ensues from crossing much-modified forms or species, we have a parallel and double series of facts, which apparently stand in close relation to each other.
It is notorious that many animals, though perfectly tamed, refuse to breed in captivity. Isidore Geoffroy St.-Hilaire[9] consequently has drawn a broad distinction between tamed animals which will not breed under captivity, and truly domesticated animals which breed freely—generally more freely, as shown in the sixteenth chapter, than in a state of nature. It is possible and generally easy to tame most animals; but experience has shown that it is difficult to get them to breed regularly, or even at all. I shall discuss this subject in detail; but will give only those cases which seem most illustrative. My materials are derived from notices scattered through various works, and especially from a Report, kindly drawn up for me by the officers of the Zoological Society of London, which has especial value, as it records all the cases, during nine years from 1838-46, in which the animals were seen to couple but produced no offspring, as well as the cases in which they never, as far as known, coupled. This MS. Report I have corrected by the annual Reports subsequently published up to the year 1865.[10] Many facts are given on the breeding of the animals in that magnificent work, ‘Gleanings from the Menageries of Knowsley Hall’ by Dr. Gray. I made, also, particular inquiries from the experienced keeper of the birds in the old Surrey Zoological Gardens. I should premise that a slight change in the treatment of animals sometimes makes a great difference in their fertility; and it is probable that the results observed in different menageries would differ. Indeed, some animals in our Zoological Gardens have become more productive since the year 1846. It is, also, manifest from F. Cuvier’s account of the Jardin des Plantes[11] that the animals formerly bred much less freely there than with us; for instance, in the Duck tribe, which is highly prolific, only one species had at that period produced young.
The most remarkable cases, however, are afforded by animals kept in their native country, which, though perfectly tamed, quite healthy, and allowed some freedom, are absolutely incapable of breeding. Rengger,[12] who in Paraguay particularly attended to this subject, specifies six quadrupeds in this condition; and he mentions two or three others which most rarely breed. Mr. Bates, in his admirable work on the Amazons, strongly insists on similar cases;[13] and he remarks, that the fact of thoroughly tamed native mammals and birds not breeding when kept by the Indians, cannot be wholly accounted for by their negligence or indifference, for the turkey and fowl are kept and bred by various remote tribes. In almost every part of the world—for instance, in the interior of Africa, and in several of the Polynesian islands—the natives are extremely fond of taming the indigenous quadrupeds and birds; but they rarely or never succeed in getting them to breed.
The most notorious case of an animal not breeding in captivity is that of the elephant. Elephants are kept in large numbers in their native Indian home, live to old age, and are vigorous enough for the severest labour; yet, with a very few exceptions, they have never been known even to couple, though both males and females have their proper periodical seasons. If, however, we proceed a little eastward to Ava, we hear from Mr. Crawfurd[14] that their “breeding in the domestic state, or at least in the half-domestic state in which the female elephants are generally kept, is of everyday occurrence;” and Mr. Crawfurd informs me that he believes that the difference must be attributed solely to the females being allowed to roam the forest with some degree of freedom. The captive rhinoceros, on the other hand, seems from Bishop Heber’s account[15] to breed in India far more readily than the elephant. Four wild species of the horse genus have bred in Europe, though here exposed to a great change in their natural habits of life; but the species have generally been crossed one with another. Most of the members of the pig family breed readily in our menageries; even the Red River hog (_Potamochœrus penicillatus_), from the sweltering plains of West Africa, has bred twice in the Zoological Gardens. Here also the Peccary (_Dicotyles torquatus_) has bred several times; but another species, the _D. labiatus,_ though rendered so tame as to be half-domesticated, is said to breed so rarely in its native country of Paraguay, that according to Rengger[16] the fact requires confirmation. Mr. Bates remarks that the tapir, though often kept tame in Amazonia by the Indians, never breeds.
Ruminants generally breed quite freely in England, though brought from widely different climates, as may be seen in the Annual Reports of the Zoological Gardens, and in the Gleanings from Lord Derby’s menagerie. The Carnivora, with the exception of the Plantigrade division, breed (though with capricious exceptions) about half as freely as ruminants. Many species of Felidae have bred in various menageries, although imported from diverse climates and closely confined. Mr. Bartlett, the present superintendent of the Zoological Gardens[17] remarks that the lion appears to breed more frequently and to bring forth more young at a birth than any other species of the family. He adds that the tiger has rarely bred; “but there are several well-authenticated instances of the female tiger breeding with the lion.” Strange as the fact may appear, many animals under confinement unite with distinct species and produce hybrids quite as freely as, or even more freely than, with their own species. On inquiring from Dr. Falconer and others, it appears that the tiger when confined in India does not breed, though it has been known to couple.
The chetah (_Felis jubata_) has never been known by Mr. Bartlett to breed in England, but it has bred at Frankfort; nor does it breed in India, where it is kept in large numbers for hunting; but no pains would be taken to make them breed, as only those animals which have hunted for themselves in a state of nature are serviceable and worth training.[18] According to Rengger, two species of wild cats in Paraguay, though thoroughly tamed, have never bred. Although so many of the Felidae breed readily in the Zoological Gardens, yet conception by no means always follows union: in the nine-year Report, various species are specified which were observed to couple seventy-three times, and no doubt this must have passed many times unnoticed; yet from the seventy- three unions only fifteen births ensued. The Carnivora in the Zoological Gardens were formerly less freely exposed to the air and cold than at present, and this change of treatment, as I was assured by the former superintendent, Mr. Miller, greatly increased their fertility. Mr. Bartlett, and there cannot be a more capable judge, says, “it is remarkable that lions breed more freely in travelling collections than in the Zoological Gardens; probably the constant excitement and irritation produced by moving from place to place, or change of air, may have considerable influence in the matter.”
Many members of the Dog family breed readily when confined. The Dhole is one of the most untamable animals in India, yet a pair kept there by Dr. Falconer produced young. Foxes, on the other hand, rarely breed, and I have never heard of such an occurrence with the European fox: the silver fox of North America (_Canis argentatus_), however, has bred several times in the Zoological Gardens. Even the otter has bred there. Every one knows how readily the semi-domesticated ferret breeds, though shut up in miserably small cages; but other species of Viverra and Paradoxurus absolutely refuse to breed in the Zoological Gardens. The Genetta has bred both here and in the Jardin des Plantes, and produced hybrids. The _Herpestes fasciatus_ has likewise bred; but I was formerly assured that the _H. griseus,_ though many were kept in the Gardens, never bred.
The Plantigrade Carnivora breed under confinement much less freely than other Carnivora, although no reason can be assigned for this fact. In the nine-year Report it is stated that the bears had been seen in the Zoological Gardens to couple freely, but previously to 1848 had most rarely conceived. In the Reports published since this date three species have produced young (hybrids in one case), and, wonderful to relate, the white Polar bear has produced young. The badger (_Meles taxus_) has bred several times in the Gardens; but I have not heard of this occurring elsewhere in England, and the event must be very rare, for an instance in Germany has been thought worth recording.[19] In Paraguay the native Nasua, though kept in pairs during many years and perfectly tamed, has never been known, according to Rengger, to breed or show any sexual passion; nor, as I hear from Mr. Bates, does this animal, or the Cercoleptes, breed in Amazonia. Two other plantigrade genera, Procyon and Gulo, though often kept tame in Paraguay, never breed there. In the Zoological Gardens species of Nasua and Procyon have been seen to couple; but they did not produce young.
As domesticated rabbits, guinea-pigs, and white mice breed so abundantly when closely confined under various climates, it might have been thought that most other members of the Rodent order would have bred in captivity, but this is not the case. It deserves notice, as showing how the capacity to breed sometimes goes by affinity, that the one native rodent of Paraguay, which there breeds _freely_ and has yielded successive generations, is the _Cavia aperea_; and this animal is so closely allied to the guinea-pig, that it has been erroneously thought to be the parent form.[20] In the Zoological Gardens, some rodents have coupled, but have never produced young; some have neither coupled nor bred; but a few have bred, as the porcupine more than once, the Barbary mouse, lemming, chinchilla, and agouti (_Dasyprocta aguti_) several times. This latter animal has also produced young in Paraguay, though they were born dead and ill-formed; but in Amazonia, according to Mr. Bates, it never breeds, though often kept tame about the houses. Nor does the paca (_Cœlogenys paca_) breed there. The common hare when confined has, I believe, never bred in Europe; though, according to a recent statement, it has crossed with the rabbit.[21] I have never heard of the dormouse breeding in confinement.
But squirrels offer a more curious case: with one exception, no species has bred in the Zoological Gardens, yet as many as fourteen individuals of _S. palmarum_ were kept together during several years. The _S. cinera_ has been seen to couple, but it did not produce young; nor has this species, when rendered extremely tame in its native country, North America, been ever known to breed.[22] At Lord Derby’s menagerie squirrels of many kinds were kept in numbers, but Mr. Thompson, the superintendent, told me that none had ever bred there, or elsewhere as far as he knew. I have never heard of the English squirrel breeding in confinement. But the species which has bred more than once in the Zoological Gardens is the one which perhaps might have been least expected, namely, the flying squirrel (_Sciuropterus volucella_): it has, also, bred several times near Birmingham; but the female never produced more than two young at a birth, whereas in its native American home she bears from three to six young.[23]
Monkeys, in the nine-year Report from the Zoological Gardens, are stated to unite most freely, but during this period, though many individuals were kept, there were only seven births. I have heard of only one American monkey, the Ouistiti, breeding in Europe.[24] A Macacus, according to Flourens, bred in Paris; and more than one species of this genus has produced young in London, especially the _Macacus rhesus,_ which everywhere shows a special capacity to breed under confinement. Hybrids have been produced both in Paris and London from this same genus. The Arabian baboon, or _ Cynocephalus hamadryas,_[25] and a Cercopithecus have bred in the Zoological Gardens, and the latter species at the Duke of Northumberland’s. Several members of the family of Lemurs have produced hybrids in the Zoological Gardens. It is much more remarkable that monkeys very rarely breed when confined in their native country; thus the Cay (_Cebus azaræ_) is frequently and completely tamed in Paraguay, but Rengger[26] says that it breeds so rarely, that he never saw more than two females which had produced young. A similar observation has been made with respect to the monkeys which are frequently tamed by the aborigines in Brazil.[27] In Amazonia, these animals are so often kept in a tame state, that Mr. Bates in walking through the streets of Para counted thirteen species; but, as he asserts, they have never been known to breed in captivity.[28]
_Birds._ Birds offer in some respects better evidence than quadrupeds, from their breeding more rapidly and being kept in greater numbers.[29] We have seen that carnivorous animals are more fertile under confinement than most other mammals. The reverse holds good with carnivorous birds. It is said[30] that as many as eighteen species have been used in Europe for hawking, and several others in Persia and India;[31] they have been kept in their native country in the finest condition, and have been flown during six, eight, or nine years;[32] yet there is no record of their having ever produced young. As these birds were formerly caught whilst young, at great expense, being imported from Iceland, Norway, and Sweden, there can be little doubt that, if possible, they would have been propagated. In the Jardin des Plantes, no bird of prey has been known to couple.[33] No hawk, vulture, or owl has ever produced fertile eggs in the Zoological Gardens, or in the old Surrey Gardens, with the exception, in the former place on one occasion, of a condor and a kite (_Milvus niger_). Yet several species, namely, the _Aquila fusca, Haliaetus leucocephalus, Falco tinnunculus, F. subbuteo,_ and _Buteo vulgaris,_ have been seen to couple in the Zoological Gardens. Mr. Morris[34] mentions as a unique fact that a kestrel (_Falco tinnunculus_) bred in an aviary. The one kind of owl which has been known to couple in the Zoological Gardens was the Eagle Owl (_Bubo maximus_); and this species shows a special inclination to breed in captivity; for a pair at Arundel Castle, kept more nearly in a state of nature “than ever fell to the lot of an animal deprived of its liberty,”[35] actually reared their young. Mr. Gurney has given another instance of this same owl breeding in confinement; and he records the case of a second species of owl, the _Strix passerina,_ breeding in captivity.[36]
Of the smaller graminivorous birds, many kinds have been kept tame in their native countries, and have lived long; yet, as the highest authority on cage-birds[37] remarks, their propagation is “uncommonly difficult.” The canary-bird shows that there is no inherent difficulty in these birds breeding freely in confinement; and Audubon says[38] that the _ Fringilla (Spiza) ciris_ of North America breeds as perfectly as the canary. The difficulty with the many finches which have been kept in confinement is all the more remarkable as more than a dozen species could be named which have yielded hybrids with the canary; but hardly any of these, with the exception of the siskin (_Fringilla spinus_), have reproduced their own kind. Even the bullfinch (_Loxia pyrrhula_) has bred as frequently with the canary, though belonging to a distinct genus, as with its own species.[39] With respect to the skylark (_Alauda arvensis_), I have heard of birds living for seven years in an aviary, which never produced young; and a great London bird-fancier assured me that he had never known an instance of their breeding; nevertheless one case has been recorded.[40] In the nine-year Report from the Zoological Society, twenty-four insessorial species are enumerated which had not bred, and of these only four were known to have coupled.
Text from Project Gutenberg, public domain in the United States.